LECTURE I
INTRODUCTORY
Everyone knows in a general way what is meant by the doctrine of descent—that it is the theory which maintains that the forms of life, animals and plants, which we see on our earth to-day, have not been the same from all time, but have been developed, by a process of transformation, from others of an earlier age, and are in fact descended from ancestors specifically different. According to this doctrine of descent, the whole diversity of animals and plants owes its origin to a transformation process, in the course of which the earliest inhabitants of our earth, extremely simple forms of life, were in part evolved in the course of time into forms of continually increasing complexity of structure and efficiency of function, somewhat in the same way as we can see every day, when any higher animal is developed from a single cell, the egg-cell, not suddenly or directly, but connected with its origin by a long series of ever more complex transformation stages, each of which is the preparation for, and leads on to the succeeding one. The theory of descent is thus a theory of development or evolution. It does not merely, as earlier science did, take for granted and describe existing forms of life, but regards them as having become what they are through a process of evolution, and it seeks to investigate the stages of this process, and to discover the impelling forces that lie behind it. Briefly, the theory of descent is an attempt at a scientific interpretation of the origin and diversity of the animate world.
In these lectures, therefore, we have not merely to show on what grounds we make this postulate of an evolution process, and to marshall the facts which necessitate it; we must also try to penetrate as far as possible towards the causes which bring about such transformations. For this reason we are forced to go beyond the limits of the theory of descent in the narrow sense, and to deal with the general processes of life itself, especially with reproduction and the closely associated problem of heredity. The transformation of species can only be interpreted in one of two ways; either it depends on a peculiar internal force, which is usually only latent in the organism, but from time to time becomes active, and then, to acertain extent, moulds it into new forms; or it depends on the continually operating forces which make up life, and on the way in which these are influenced by changing external conditions. Which of these alternatives is correct we can only undertake to determine when we know the phenomena of life, and as far as possible their causes, so that it is indispensable to make ourselves acquainted with these as far as we can.
When we look at one of the lowest forms of life, such as an Amœba or a single-celled Alga, and reflect that, according to the theory of evolution, the whole realm of creation as we see it now, with Man at its head, has evolved from similar or perhaps even smaller and simpler organisms, it seems at first sight a monstrous assumption, and one which quite contradicts our simplest and most certain observations. For what is more certain than that the animals and plants around us remain the same, as long as we can observe them, not through the lifetime of an individual only, but through centuries, and in the case of many species, for several thousand years?
This being so, it is intelligible enough that the doctrine of evolution, on its first emergence at the end of the eighteenth century, was received with violent opposition, not on the part of the laity only, but by the majority of scientific minds, and instead of being followed up, was at first opposed, then neglected, and finally totally forgotten, to spring up anew in our own day. But even then a host of antagonists ranged themselves against the doctrine, and, not content with loftily ignoring it, made it the subject of the most violent and varied attacks.
This was the state of affairs when, in 1858, Darwin's book onThe Origin of Speciesappeared, and hoisted the flag of evolution afresh. The struggle that ensued may now be regarded as at an end, at least as far as we are concerned—that is, in the domain of science. The doctrine of descent has gained the day, and we can confidently say that the Evolution theory has become a permanent possession of science that can never again be taken away. It forms the foundation of all our theories of the organic world, and all further progress must start from this basis.
In the course of these lectures, we shall find at every step fresh evidence of the truth of this assertion, which may at first seem all too bold. It is not by any means to be supposed that the whole question in regard to the transformation of organisms and the succession of new forms of life has been answered in full, or that we have now been fortunate enough to solve the riddle of life itself. No! whether we ever reach that goal or not, we are a long way from it as yet, andeven the much easier problem, how and by what forces the evolution of the living world has proceeded from a given beginning, is far from being finally settled; antagonistic views are still in conflict, and there is no arbitrator whose authoritative word can decide which is right. TheHow?of evolution is still doubtful, but not thefact, and this is the secure foundation on which we stand to-day: The world of life, as we know it, has been evolved, and did not originate all at once.
Were I to try to give, in advance, even an approximate idea of the confidence with which we can take our stand on this foundation, I should be almost embarrassed by the wealth of facts on which I might draw. It is hardly possible nowadays to open a book on the minute or general structural relations, or on the development of any animal whatever, without finding in it evidences in favour of the Evolution theory, that is to say, facts which can only be understood on the assumption of the evolution of the organic world. This, too, without taking into account at all the continually increasing number of facts Palæontology is bringing to light, placing before our eyes the forms which the Evolution theory postulates as the ancestors of the organic world of to-day: birds with teeth in their bills, reptile-like forms clothed with feathers, and numerous other long-extinct forms of life, which, covered up by the mud of earlier waters, and preserved as 'fossils' in the later sedimentary rocks, tell us plainly how the earlier world of animals and plants was constituted. Later, we shall see that the geographical distribution of plant and animal species of the present day can only be understood in the light of the Evolution theory. But meantime, before we go into details, what may justify my assumption is the fact that the Evolution theory enables us to predict.
Let us take only a few examples. The skeleton of the wrist in all vertebrate animals above Fishes consists of two rows of small bones, on the outer of which are placed the five bones of the palm, corresponding to the five fingers. The outer row is curved, and there is thus a space between the two rows, which, in Amphibians and Reptiles, is filled by a special small bone. This 'os centrale' is absent in many Mammals, notably, for instance, in Man, and the space between the two rows is filled up by an enlargement of one of the other bones. Now if Mammals be descended from the lower vertebrates, as the theory of descent assumes, we should expect to find the 'os centrale' even in Man in young stages, and, after many unsuccessful attempts, Rosenberg has at last been able to demonstrate it at a very early stage of embryonic development.
This prediction, with another to be explained later, is based upon the experience that the development of an individual animal follows, in a general way, the same course as the racial evolution of the species, so that structures of the ancestors of a species, even if they are not found in the fully developed animal, may occur in one of its earlier embryonic stages. Further on, we shall come to know this fact more intimately as a 'biogenetic law,' and it alone would be almost enough to justify the theory of evolution. Thus, for instance, the lowest vertebrates, the Fishes, breathe by means of gills, and these breathing organs are supported by four or more gill-arches, between which spaces, the gill-slits, remain open for the passage of water. Although Reptiles, Birds, and Mammals breathe by lungs, and at no time of their life by gills, yet, in their earliest youth, that is, during their early development in the egg, they possess these gill-arches and gill-slits, which subsequently disappear, or are transformed into other structures.
On the strength of this 'biogenetic law' it could also be predicted that Man, in whom, as is well known, there are twelve pairs of ribs, would, in his earliest youth, possess a thirteenth pair, for the lower Mammals have more numerous ribs, and even our nearest relatives, the anthropoid Apes, the gorilla and chimpanzee, have a thirteenth rib, though a very small one, and the siamang has even a fourteenth. This prediction also has been verified by the examination of young human embryos, in which a small thirteenth rib is present, though it rapidly disappears.
During the seventies I was engaged in investigating the development of the curious marking which adorns the long body of many of our caterpillars. I studied in particular the caterpillars of our Sphingidæ or hawk-moths, and found, by a comparison of the various stages of development from the emergence of the caterpillar from the egg on to its full growth, that there is a definite succession of different kinds of markings following each other, in a whole range of species, in a similar manner. From the standpoint of the Evolution theory, I concluded that the markings of the youngest caterpillars, simple longitudinal stripes, must have been those of the most remote ancestors of our present species, while those of the later stages, oblique stripes, were those of ancestors of a later date.
If this were the case, then all the species of caterpillar which now exhibit oblique stripes in their full-grown stage must have had longitudinal stripes in their youngest stages, and because of this succession of markings in the individual development, I was able to predict that the then unknown young form of the caterpillar of ourprivet hawk-moth (Sphinx ligustri) must have a white line along each side of the back. Ten years later, the English zoologist, Poulton, succeeded in rearing the eggs ofSphinx ligustri, and it was then demonstrated that the young caterpillar actually possessed the postulated white lines.
Such predictions undoubtedly give the hypothesis on which they are based, the Evolution theory, a high degree of certainty, and are almost comparable to the prediction of the discovery of the planet Neptune by Leverrier. As is well known, this, the most distant of all the planets, whose period of revolution round the sun is almost 165 of our years, would probably never have been recognized as a planet, had not Adams, an astronomer at the Greenwich Observatory, and afterwards Leverrier, deduced its presence from slight disturbances in the path of Jupiter's moons, and indicated the spot where an unknown planet must be looked for. Immediately all telescopes were directed towards the spot indicated, and Galle, at the Berlin Observatory, found the sought-for planet.
We might with justice regard as lacking in discernment those who, in the face of such experiences, still doubt that the earth revolves round the sun, and we might fairly say the same of any one who, in the face of the known facts, would dispute the truth of the Evolution theory. It is the only basis on which an understanding of these facts is possible, just as the Kant-Laplace theory of the solar system is the only basis on which an adequate interpretation of the facts of the heavens can be arrived at.
To this comparison of the two theories it has been objected that the Evolution theory has far less validity than the other, first, because it can never be mathematically demonstrated, and secondly, because at the best it can only interpret the transformations of the animate world, and not its origin. Both objections are just: the phenomena of life are in their nature much too intricate for mathematics to deal with, except with extreme diffidence; and the question of the origin of life is a problem which will probably have to wait long for solution. So, if it gives pleasure to any one to regard the one theory as having more validity than the other, no one can object; but there is no particular advantage to be gained by doing so. In any case, the Evolution theory shares the disadvantage of not being able to explain everything in its own province with the Kant-Laplace cosmogony, for that, too, must presuppose the first beginning, the rotating nebula.
Although I regard the doctrine of descent as proved, and hold it to be one of the greatest acquisitions of human knowledge,I must repeat that I do not mean to say that everything is clear in regard to the evolution of the living world. On the contrary, I believe that we still stand merely on the threshold of investigation, and that our insight into the mighty process of evolution, which has brought about the endless diversity of life upon our earth, is still very incomplete in relation to what may yet be found out, and that, instead of being vainglorious, our attitude should be one of modesty. We may well rejoice over the great step forward which the dominant recognition of the Evolution theory implies, but we must confess that the beginnings of life are as little clear to us as those of the solar system. But we can do this at least: we can refer the innumerable and wonderful inter-relations of the organic cosmos to their causes—common descent and adaptation—and we can try to discover the ways and means which have co-operated to bring the organic world to the state in which we know it.
When I say that the theory of descent is the most progressive step that has yet been taken in the development of human knowledge, I am bound to give my reasons for this opinion. It is justified, it seems to me, even by this fact alone, that the Evolution idea is not merely a new light on the special region of biological science, zoology and botany, but is of quite general importance. The conception of an evolution of the world of life upon the earth reaches far beyond the bounds of any single science, and influences our whole realm of thought. It means nothing less than the elimination of the miraculous from our knowledge of nature, and the placing of the phenomena of life on the same plane as the other natural processes, that is, as having been brought about by the same forces, and being subject to the same laws. In the domain of the inorganic, no one now doubts that out of nothing nothing can come: energy and matter are from everlasting to everlasting, they can neither be increased or decreased, they can only be transformed—heat into mechanical energy, into light, into electricity, and so on. For us moderns, the lightning is no longer hurled by the Thunderer Zeus on the head of the wicked, but, careless alike of merit or guilt, it strikes where the electric tension finds the easiest and shortest line of discharge. Thus to our mode of thought it now seems clear that no event in the world of the living depends upon caprice, that at no time have organisms been called forth out of nothing by the mighty word of a Creator, but they have been produced at all times by the co-operation of the existing forces of nature, and every species must have arisen just where, and when, and in the form in which it actually did arise, as the necessary outcome of the existing conditions of energy andmatter, and of their interactions upon each other. It is this correlation of animate nature with natural forces and natural laws which gives to the doctrine of evolution its most general importance. For it thus supplies the keystone in the arch of our interpretation of nature and gives it unity; for the first time it makes it possible to form a conception of a world-mechanism, in which each stage is the result of the one before it, and the cause of the succeeding one.
How deeply all our earlier opinions are affected by this doctrine will become clear if we fix our attention on a single point, the derivation of the human understanding from that of animal ancestors. What of the reason of Man, of his morals, of his freedom of will? may be asked, as it has been, and still is often asked. What has been regarded as absolutely distinct from the nature of animals is said to differ from their mental activities only in degree, and to have evolved from them. The mind of a Kant, of a Laplace, of a Darwin—or to ascend into the plane of the highest and finest emotional life, the genius of a Raphael or a Mozart—to have any real connexion, however far back, with the lowly psychical life of an animal! That is contrary to all our traditionary, we might say our inborn, ideas, and it is not to be wondered at that the laity, and especially the more cultured among them, should have opposed such a doctrine whose dominating power was unintelligible to them, because they were ignorant of the facts on which it rests. That a man should feel his dignity lowered by the idea of descent from animals is almost comical to the naturalist, for he knows that every one of us, in his first beginning, occupied a much lowlier position than that of our mammalian ancestors—was, in fact, as regards visible structure, on a level with the Amœba, that microscopically minute unicellular animal, which can hardly be said to possess organs, and whose psychical activities are limited to recognizing and engulfing its food. Very gradually at first, and step by step, there develop from this single cell, the ovum, more and more numerous cells; this mass of cells segregates into different groups, which differentiate further and further, until at last they form the perfect man. This occurs in the development of every human being, and we are merely unaccustomed to the thought that it means nothing else than an incredibly rapid ascent of the organism from a very low level of life to the highest.
Still less is it to be wondered at that the Evolution doctrine met with violent opposition on the part of the representatives of religion, for it stood in open contradiction to that remarkable and venerable cosmogony, the Mosaic story of Creation, which people had been accustomed to regard, not as what it is—a conception of natureat an early stage of human culture—but as an inalienable part of our own religion. But investigation shows us that the doctrine of evolution is true, and it is only a weak religion which is incapable of adapting itself to the truth, retaining what is essential, and letting go what is unessential and subject to change with the development of the human mind. Even the heliocentric hypothesis was in its day declared false by the Church, and Galilei was forced to retract; but the earth continued to revolve round the sun, and nowadays any one who doubted it would be considered mentally weak or warped. So in all likelihood the time is not far distant when the champions of religion will abandon their profitless struggle against the new truth, and will see that the recognition of a law-governed evolution of the organic world is no more prejudicial to true religion than is the revolution of the earth round the sun.
Having given this very general orientation of the Evolution problem, which is to engage our attention in detail, I shall approach the problem itself by the historical method, for I do not wish to bring the views of present-day science quite suddenly and directly into prominence. I would rather seek first to illustrate how earlier generations have tried to solve the question of the origin of the living world. We shall see that few attempts at solution were made until quite recently, that is, until the end of the eighteenth and the beginning of the nineteenth century. Only then there appeared a few gifted naturalists with evolutionist ideas, but these ideas did not penetrate far; and it was not till after the middle of the nineteenth century that they found a new champion, who was to make them common property and a permanent possession of science. It was the teaching of Charles Darwin that brought about this thorough awakening, and laid the foundations of our present interpretations, and his work will therefore engross our attention for a number of lectures. Only after we have made ourselves acquainted with his teaching shall we try to test its foundations, and to see how far this splendid structure stands on a secure basis of fact, and how deeply its power of interpretation penetrates towards the roots of phenomena. We shall examine the forces by which organisms are dominated, and the phenomena produced, and thereby test Darwin's principles of interpretation, in part rejecting them, in part accepting them, though in a much extended form, and thus try to give the whole theoretic structure a more secure foundation. I hope to be able to show that we have made some real progress since Darwin's day, thatdeductions have been drawn from his theory which even he did not dream of, which have thrown fresh light on a vast range of phenomena, and, finally, that through the more extended use of his own principles, the Evolution theory has gained a completeness, and an intrinsic harmony which it previously lacked.
This at least is my own opinion, but I cannot ignore the fact that it is by no means shared by all living naturalists. The obvious gaps and insufficiencies of the Darwinian theory have in the last few decennia prompted all sorts of attempts at improving it. Some of these were lost sight of almost as soon as they were suggested, but others have held their own, and can still claim numerous supporters. It would only tend to bewilder if I gave an account of those of the former class, but those which still hold their own must be noticed in these lectures, though it is by no means my intention to expound the confused mass of opinions which has gathered round the doctrine of evolution, but rather to give a presentation of the theory as it has gradually grown up in my own mind in the course of the last four decades. Even this will not be the last of which science will take knowledge, but it will, I hope, at least be one which can be further built upon.
Let us, then, begin at once with that earliest forerunner of the modern theory of descent, the gifted Greek philosopher Empedocles, who, equally important as a leader of the state of Agrigentum, and as a thinker in purely theoretical regions of thought, advanced very notable views regarding the origin of organisms. We must, however, be prepared to hear something that is hardly a theory in the modern scientific acceptation of that term; and we must not be repelled by the unbridled poetical fancy of the speculative philosopher; we have to recognize that there is a sound kernel contained in his amusing pictures—a thought which we meet with later, in much more concrete form, in the Darwinian theory, and which, if I mistake not, we shall keep firm hold of in all time to come.
According to Empedocles the world was formed by the four elements of the ancients, Earth, Water, Fire, and Air, moved and guided by two fundamental forces, Hate and Love, or, as we should now say, Repulsion and Attraction. Through the chance play of these two forces with the elements, there arose first the plants, then the animals, in such a manner that at first only parts and organs of animals were formed: single eyes without faces, arms without bodies, and so on. Then, in wild play, Nature attempted to put together these separate parts, and so created all manner of combinations, for the most part inept monsters unfit for life, but in a fewcases, where the parts fitted, there resulted a creature capable not only of life, but, if the juxtaposition was perfect, even of reproduction.
This phantastic picture of creation seems to us mad enough, but there slumbers in it, all unsuspected though it may have been by the author, the true idea of selection, the idea that much that is unfit certainly arises, but that only the fit endures. The mechanical coming-to-be of the fit is the sound kernel in this wondersome doctrine.
The natural science of the ancients, in regard to life and its forms, reached its climax in Aristotle (died 322B. C.). A true polyhistorian, his writings comprehended all the knowledge of his time, but he also added much to it from his own observation. In his writings we find many good observations on the structure and habits of a number of organisms, and he also had the merit of being the first to attempt a systematic grouping of animals. With true insight, he grouped all the vertebrates together as Enaimata or animals with blood, and classed all the rest together as Anaimata or bloodless animals. That he denied to the latter group the possession of blood is not to be wondered at, when we take into account the extremely imperfect means of investigation available in his time, nor is it surprising that he should have ranked this motley company, in antithesis to the blood-possessing animals, as a unified and equivalent group. Two thousand years later, Lamarck did exactly the same thing, when he divided the animals into backboned and backboneless, and we reckon this nowadays as a merit only in so far that he was the first, after Aristotle, to re-express the solidarity of the classes of animals which we now call vertebrates.
Aristotle was, however, not a systematic zoologist in our sense of the term, as indeed was hardly possible, considering the very small number of animal forms that were known in his time. In our day we have before us descriptions of nearly 300,000 named species wherefrom to construct our classification, while Aristotle knew hardly more than 200. Of the whole world of microscopic animals he could, of course, have no idea, any more than of the remains of prehistoric animals, of which we now know about 40,000 named and adequately described species. One would have thought that it would have occurred to a quick-witted people like the Greeks to pause and ponder when they found mussel-shells and marine snail-shells on the hills far above the sea; but they explained these by the great flood in the time of Deucalion and Pyrrha, and they did not observe that the fossil molluscs were of different species from the similar animals living in the sea in their own day.
Thus there was nothing to suggest to Aristotle and others of his time the idea that a transformation of species had been going on through the ages, and even the centuries after him evoked no such idea, nor did there arise new speculations, after the manner of Empedocles, in regard to the origin of the organic world. On the whole, the knowledge of the living world retrograded rather than advanced until the beginning of the Roman Empire. What Aristotle had known was forgotten, and Pliny's work on animals is a catalogue embellished with numerous fables, arranged according to a purely external principle of division. Pliny divided animals into those belonging to earth, water, and air, which is not very much more scientific than if he had arranged them according to the letters of the alphabet.
During the time of the Roman Empire, as is well known, the knowledge of natural history sank lower and lower; there was no more investigation of nature, and even the physicians lost all scientific basis, and practised only in accordance with their traditional esoteric secrets. As the whole culture of the West gradually disappeared, the knowledge of nature possessed by earlier centuries was also completely lost, and in the first half of the Middle Ages Europeans revealed a depth of ignorance of the natural objects lying about them, which it is difficult for us now to form any conception of.
Christianity was in part responsible for this, because it regarded natural science as a product of heathendom, and therefore felt bound to look coldly on it, if not even to oppose it. Later, however, even the Christian Church felt itself forced to give the people some mental nourishment in the form of natural history, and under its influence, perhaps actually composed by teachers of the Church, there appeared a little book, the so-calledPhysiologus, which was meant to instruct the people in regard to the animal world. This remarkable work, which has been preserved, must have had a very wide distribution in the earlier Middle Ages, for it was translated into no fewer than twelve languages, Greek, Armenian, Syriac, Arabic, Ethiopic, and so on. The contents are very remarkable, and come from the most diverse sources, that is, from the most different writers of antiquity, from Herodotus, from the Bible, and so forth, but never from original observation. The compilation does not really give descriptions of animals or of their habits, but, of each of the forty-one animals which thePhysiologusrecognizes, something remarkable is briefly related in true lapidary style, sometimes a mere curiosity without further import, or sometimes a symbolical interpretation. Thus the book says of the panther: 'he is gaily coloured; after satiating himself hesleeps three days, and awakes roaring, giving forth such an agreeable odour that all animals come to him.' Of the pelican the well-known legend is related, that it tears open its own breast to feed its young with its blood, thus standing as a symbol of mother-love. Fabulous creatures, too, appear in these pages. Of the Phœnix, that bird whose plumage glitters with gold and precious stones, which was known even to Herodotus, and which has survived through Eastern fairy-tales on to the time of our own romanticists (Tieck), we read: 'it lives a thousand years, because it has not eaten of the tree of knowledge'; then 'it sets fire to itself and arises anew from its own ashes,' a symbol of nature's infinite power of renewing its youth.
But while among the peoples of Europe all the science of the ancients was lost, except a few barely recognizable fragments, the old lore was preserved, both as regards organic nature and other orders of facts, among the Arabs, through whom so many treasures of antiquity have eventually been handed down to us, coming in the track of the Arabian conquests across North Africa and Spain to the nations of Europe.
It was in this way, too, that the writings of Aristotle again found recognition, after having been translated into Latin at Palermo at the order of that enthusiast for Science and Art, the Hohenstaufen Emperor, Frederick the Second. Our Emperor presented one copy of Aristotle's writings to the University of Bologna, and thus the wisdom of the ancient Greeks again became the common property of European culture. From the thirteenth century to the eighteenth, the study of natural science was limited to repeating and extending the work of Aristotle. Nothing new, depending upon personal observation, was added, and it does not even seem to have occurred to any one to subject the statements of the Stagirite to any test, even when they concerned the most familiar objects. No one noticed the error which ascribed to the fly eight legs instead of six; there was in fact as yet no investigation, and all knowledge of natural history was purely scholastic, and gave absolute credence to the authority of the ancients.
A revulsion, however, occurred in the century of the Reformation, with the breaking down of the blind belief in authority which had till then prevailed in all provinces of human knowledge and thought. After a long and severe struggle, dry scholasticism was finally overcome, and natural science, with the rest, turned from a mere reliance on books to original thinking and personal observation. Thenceforward interpretations of natural processes were sought for no longer in the writings of the ancients, but in Nature herself.Of the magnitude of this emancipation, and of the severity of the struggle against deep-rooted authority, one could form a faint idea from experience even in my own youth. Our young minds were so deeply imbued with the involuntary feeling that the ancients were superior to us moderns in each and every respect, that not only the hardly re-attainable plastic art of the Greeks and the immortal songs of Homer, but all the mental products of antiquity seemed to us models which could never be equalled; the tragedies of Sophocles were for us the greatest tragedies that the world had ever seen, the odes of Horace the most beautiful poems of all time!
In the domain of natural science the new era began with the overthrow of the Ptolemaic cosmogony, which, for more than a thousand years, had served as a basis for astronomy. When the German canon, Nicolas Copernicus (born at Thorn, 1473, died 1543), reversed the old theory, and showed that the sun did not revolve round the earth, but the earth round the sun, the ice was broken and the way paved for further progress. Galilei uttered his famous 'e pur si muove,' Kepler established his three laws of the movements of the planets, and Newton, a century later, interpreted their courses in terms of the law of gravitation.
But we have not here to do with a history of physics or astronomy, and I only wish to recall these well-known facts, in order that we may see how increased knowledge in this domain was always accompanied by advances in that of biology.
Here, however, we cannot yet chronicle any such thoroughgoing revolution of general conceptions; the basis of detailed empirical knowledge was not nearly broad enough for that, and it was in the acquiring of such a foundation that the next three centuries, from the sixteenth to the end of the eighteenth, were eagerly occupied.
The first step necessary was to collate the items of individual knowledge in regard to the various forms of life, and to bring the whole in unified form into general notice. This need was met for the first time by Conrad Gessner'sThierbuch, a handsome folio volume, printed at Zurich in 1551, and embellished with numerous woodcuts, some of them very good. This was followed, in 1600, by a great work in many volumes, written in Latin, by a professor of Bologna, Aldrovandi. Not native animals alone but foreign ones also were described in these works, for, after the discovery of America and the opening up of communication with the East Indies, many new animal and plant forms came to the knowledge of European nations by way of the sea. Thus Francesco Hernandez (died 1600), physician in ordinary to Philip II, described no fewer than forty newMammals, more than two hundred Birds, and many other American animals.
Again, in a quite different way, the naturalist's field of vision was widened, namely, by the invention of the simple microscope, with which Leeuwenhoek first discovered the new world of Infusorians, and Swammerdam made his notable observations on the structure and development of the very varied minute animal inhabitants of fresh water. In the same century, the seventeenth, anatomists like Tulpius, Malpighi, and many others extended the knowledge of the internal structure of the higher animals and of Man, and a foundation was laid for a deeper insight into the nature of vital functions by the discovery of the circulation of the blood in Man and the higher animals. In the following century, the eighteenth, this path of active research was eagerly followed, and we need only mention such names as Réaumur, Rösel von Rosenhof, De Geer, Bonnet, J. Chr. Schäfer, and Ledermüller, to be immediately reminded of the wealth of facts about the structure, life, and especially the development of our indigenous animals, which we owe to the labours of these men.
All these advances, great and many-sided as they were, did not at once lead to a renewal of the attempt of Empedocles to explain the origin of the organic world. This was as yet not even recognized as a problem requiring investigation, for men were content to take the world of life simply as a fact. The idea of getting beyond the naïve, poetic standpoint of the Mosaic story of Creation was as yet remote from the minds of naturalists, partly because they were wholly fascinated by the observation of masses of details, but chiefly because, first by the English physician, John Ray (died 1678), then by the great Swede, Carl Linné, the conception of organic 'species' had been formulated and sharply defined. It is true enough that before the works of these two men 'species' had been spoken of, but without being connected with any definite idea; the word was used rather in the same vague sense as the word 'genus,' to designate one of the smaller groups of organic forms, but without implying any clear idea of its scope or of its limitations. Now, however, for the first time, the term 'species' came to be used strictly to mean the smallest homogeneous group of individual forms of life upon the earth. John Ray held that the surest indication of a 'species' was that its members had been produced from the same seed; that is, 'forms which are of different species maintain this specific nature constantly, and one species does not arise from the seed of another.' Here we have the germ of the doctrine of the absolute nature and theimmutability of species which Linné briefly characterized in these words: 'Species tot sunt, quot formæ ab initio creatæ sunt,' 'there are just so many species as there were forms created in the beginning.' It is here clearly implied, that species as we know them have been as they are from all time, that, therefore, they exist in nature as such and unchangeably, and have not been merely read into nature by man.
This view, though we cannot now regard it as correct, was undoubtedly reasonable, and thoroughly in accordance with the spirit of the time; it was congruent with the knowledge, and above all with the scientific endeavours of the age. In the eighteenth century there was danger that all outlook on nature as a whole would be lost—smothered under the enormous mass of isolated facts, and especially under the inundation of diverse animal and plant forms which were continually being recognized. It must therefore have been regarded as a real deliverance, when Linné reduced this chaos of forms to a clearly ordered system, and relegated each form to its proper place and value in relation to the whole. How, indeed, could the great systematist have performed his task at all, if he had not been able to work with definite and sharply circumscribed groups of forms, if he had not been able to regard at least the lowest elements of his system, the species, as fixed and definite types? On the other hand, Linné was much too shrewd an observer not to entertain, in the course of his long life, and under the influence of the continually accumulating material, doubts as to the correctness of his assumption of the fixity and absoluteness of his species. He discovered from his own experience, what is fully borne out by ours, that it is easy enough to define a species when there are only a few specimens of a form to deal with, but that the difficulty increases in proportion to the number and to the diversity of habitat of those that are to be brought under one category. In the last edition of theSystema Naturæwe find very noteworthy passages in which Linné wonders whether, after all, a species may not change, and in the course of time diverge into varieties, and so forth. Of these doubts no notice was taken at the time; the accepted doctrine of the fixity of species was held to and even raised to the rank of a scientific dogma. Georges Cuvier, the great disciple of the Stuttgart 'Karlschule,' accentuated the doctrine still further by his establishment of animal-types, the largest groups of forms in the animal kingdom within which a definite and fundamentally distinct plan of architecture prevails. His four types, Vertebrates, Molluscs, Articulate and Radiate animals, furnished a further corroboration of the absolute nature of species, since theyseemed to show that even the highest and most comprehensive groups are sharply defined off from one another.
Let me add that this doctrine of the absolute nature of species was not fully elaborated till our own day, when the Swiss (afterwards American) naturalist, Louis Agassiz, went so far as to maintain that not only the highest and the lowest categories, but all those coming between them, were categories established and sharply separated by Nature herself. But in spite of much ingenuity and his wide and comprehensive outlook he exerted himself in vain to find satisfactory and really characteristic definitions of what was to be considered a class, an order, a family, or a genus. He did not succeed in finding a rational definition of these systematic concepts, and his endeavour may be regarded as the last important attempt to prop up an interpretation of nature already doomed to fall. But in referring to Louis Agassiz I have anticipated the historical course of scientific development, and must therefore go back to the last quarter of the eighteenth century.
The first unmistakable pioneer of the theory of descent, which now emerged for the first time as a scientific doctrine, was our great poet Goethe. He has indeed been often named as the founder of the theory, but that seems to me saying too much. It is true, however, that the inquiring mind of the poet certainly recognized in the structure of 'related' animals the marvellous general resemblances amid all the differences in detail, and he probed for the reason of these form-relations. Through the science of 'comparative anatomy,' as it was taught at the close of the century by Kielmeyer, Cuvier's teacher, and later by Cuvier himself, Blumenbach, and others, numerous facts had become known, which paved the way for such questions. It had, for instance, been recognized that the arm of man, the wing of the bird, the paddle of the seal, and even the foreleg of the horse, contain essentially the same chain of bones, and Goethe had already expressed these relations in his well-known verse,
'Alle Gestalten sind ähnlich, doch keine gleichet der andern,Und so deutet der Chor auf ein geheimes Gesetz.'
'Alle Gestalten sind ähnlich, doch keine gleichet der andern,Und so deutet der Chor auf ein geheimes Gesetz.'
'Alle Gestalten sind ähnlich, doch keine gleichet der andern,Und so deutet der Chor auf ein geheimes Gesetz.'
'Alle Gestalten sind ähnlich, doch keine gleichet der andern,
Und so deutet der Chor auf ein geheimes Gesetz.'
As to what this law was he did not at that time pronounce an opinion, though he may even then have thought of the transformation of species. At first he contented himself with seeking for an ideal archetype or 'Urtypus' which was supposed to lie at the foundation of a larger or smaller group. He discovered the archetypal plant or 'Urpflanze,' when he rightly recognized that the parts of the flower are nothing more than modified leaves. He spoke plainly of the 'metamorphosis of plants,' meaning by that the transformation of his 'archetype' into theendless diversity of actual plant forms. But at first he certainly thought of this transformation only in the ideal sense, and not as a factual evolutionary process.
The first who definitely maintained the latter view was, remarkably enough, the grandfather of the man who, in our own day, made the theory of descent finally triumphant, the English physician Erasmus Darwin, born 1731. This quiet thinker published, in 1794, a book entitledZoonomia, and in it he takes the important step of substituting for Goethe's 'secret law' a real relationship of species. He proclaims the gradual establishment and ennobling of the animal world, and bases his view mainly on the numerous obvious adaptations of the structure of an organ to its use. I have not been able to find any passage in the book in which he has expressly indicated that, because many of the conditions of life could not have existed from the beginning, these adaptations are therefore, as such, an argument for the gradual transformation of species. But he assumed that such exact adaptations to the functions of an organ could only arise through the exercise of that function, and in this he saw a proof of transformation. Goethe had expressed the same idea when he said, 'Thus the eagle has conformed itself through the air to the air, the mole through the earth to the earth, and the seal through the water to the water,' and this shows that he too at one time thought of an actual transformation. But neither he nor Erasmus Darwin were at all clear as tohowthe use of an organ could bring about its variation and transformation. The latter only says that, for instance, the snout of the pig has become hard through its constant grubbing in the ground; the trunk of the elephant has acquired its great mobility through the perpetual use of it for all sorts of purposes; the tongue of the herbivore owes its hard, grater-like condition to the rubbing to and fro of the hard grass in the mouth, and so on. How acute and thoughtful an observer Erasmus Darwin was, is shown by the fact that he had correctly appreciated the biological significance of many of the colour-adaptations of animals to their surroundings, though it was reserved for his grandson to make this fully clear at a much later date. Thus he regarded the varied colouring of the python, of the leopard, and of the wild cat as the best adapted for concealing them from their prey amid the play of light and shadow in a leafy thicket. The black spot in front of the eye of the swan he considered an arrangement to prevent the bird from being dazzled, as would happen if that spot were as snow-white as the rest of the plumage.
At the end of the book he sums up his views in the followingsentences: 'The world has been evolved, not created; it has arisen little by little from a small beginning, and has increased through the activity of the elemental forces embodied in itself, and so has rather grown than suddenly come into being at an almighty word.' 'What a sublime idea of the infinite might of the great Architect! the Cause of all causes, the Father of all fathers, the Ens entium! For if we could compare the Infinite it would surely require a greater Infinite to cause the causes of effects than to produce the effects themselves.'
In these words he sets forth his position in regard to religion, and does so in precisely the same terms as we may use to-day when we say: 'All that happens in the world depends on the forces that prevail in it, and results according to law; but where these forces and their substratum, Matter, come from, we know not, and here we have room for faith.'
I have not been able to discover whether theZoonomia, with its revolutionary ideas, attracted much attention at the time when it appeared, but it would seem not. In any case, it was afterwards so absolutely forgotten, that in an otherwise very completeHistory of Zoology, published in 1872 by Victor Carus, it was not even mentioned. About a year after the appearance ofZoonomia, Isidore Geoffrey St.-Hilaire in Paris expounded the view that what are called species are really only 'degenerations,' deteriorations from one and the same type, which shows that he too had begun to have doubts as to the fixity of species. Yet it was not till the third decade of the nineteenth century that he clearly and definitely took up the position of the doctrine of transformation, and to this we shall have to return later on.
But as early as the first decade of the century this position was taken up by two noteworthy naturalists, a German and a Frenchman, Treviranus and Lamarck.
Gottfried Reinhold Treviranus, born at Bremen in 1776, an excellent observer and an ingenious investigator, published, in 1802, a book entitledBiologie, oder Philosophie der lebenden Natur[Biology, or Philosophy of Animate Nature], in which he expresses and elaborates the idea of the Evolution theory with perfect clearness. We read there, for instance: 'In every living being there exists a capacity for endless diversity of form; each possesses the power of adapting its organization to the variations of the external world, and it is this power, called into activity by cosmic changes, which has enabled the simple zoophytes of the primitive world to climb to higher and higher stages of organization, and has brought endlessvariety into nature.' But where the motive power lies, which brings about these transformations from the lowliest to ever higher forms of life, was a question which Treviranus apparently did not venture to discuss. To do this, and thus to take the first step towards a causal explanation of the assumed transformations, was left for his successor.
Jean Baptiste de Lamarck, born in 1744 in a village of Picardy, was first a soldier, then a botanist, and finally a zoologist. He won his scientific spurs first by hisFlora of France, and zoology holds him in honour as the founder of the category of 'vertebrates.' Not that he occupied himself in particular detail with these, but he recognized the close alliance of the classes of animals in question—an alliance which was subsequently expressed by Cuvier by the systematic term 'type' or 'embranchement.'
In hisPhilosophie zoologique, published in 1809, Lamarck set forth a theory of evolution whose truth he attempted to vindicate by showing—as Treviranus had done before him—that the conception of species, on the immutability of which the whole hypothesis of creation had been based, was an artificial one, read into nature by us; that sharply circumscribed groups do not exist in nature at all; and that it is often very difficult, and not infrequently quite impossible, to define one species precisely from allied forms, because it is connected with these on all sides by transition stages. Groups of forms which thus melted into one another indicated that the doctrine of the fixity of species could not be correct, any more than that of their absolute nature. Species, he maintained, are not immutable, and are not so old as nature; they are fixed only for a certain time. The shortness of our life prevents our directly recognizing this. 'If we lived a much shorter time, say about a second, the hour-hand of the clock would appear to us to stand still, and even the combined observations of thirty generations would afford no decisive evidence as to the hand's movement, and yet it had been moving.'
The causes on which, according to Lamarck, the transformation of species, their modification into new species, depends, lie in the changes in the conditions of life which must have occurred ceaselessly from the earliest period of the earth's history till our own day, now here, now there, due in part to changes in climate and in food-supply, in part to changes in the earth's crust by the rising or sinking of land-masses, and so forth. These external changes have sometimes been thedirectcause of changes in bodily structure, as in the case of heat or cold; but they have sometimes and much more effectively operatedindirectly. Thus changed conditions may haveprompted an animal of a given species to use certain parts of its body in a new way, more vigorously, or less actively, or even not at all, and the more vigorous use, or, conversely, the disuse, has brought about variations in the organ in question.
Thus the whales lost their teeth when they abandoned their fish diet, and acquired the habit of feeding on minute and delicate molluscs, which they swallowed whole without seizure or mastication. Thus, too, the eyes of the mole degenerated through its life in the dark, and a still greater degeneration of the eyes has taken place in animals, like the proteus-salamander, which always inhabit lightless caves. In mussels both head and eyes degenerated because the animals could no longer use them after they became enclosed in opaque mantles and shells. In the same way snakes lost their legspari passuwith the acquisition of the habit of moving along by wriggling their long bodies, and of creeping through narrow fissures and holes. On the other hand, Lamarck interpreted the evolution of the web-feet of swimming birds by supposing that some land-bird or other had formed the habit of going into the water to seek for food, and consequently of spreading out its toes as widely as possible so as to strike the water more vigorously. In this way the fold of skin between the toes was stretched, and as the extension of the toes was very frequent and was continued through many generations, the web expanded and grew larger, and thus formed the web-foot.
In the same way the long legs of the wading birds have been, according to Lamarck, gradually evolved by the continual stretching of the limbs by wading in deeper and deeper water, and similarly for the long necks and bills of the waders, the herons and the storks. Finally we may mention the case of the giraffe, whose enormously long neck and tall forelegs are interpreted as due to the fact that the animal feeds on the foliage of trees, and was always stretching as far as possible, in order to reach the higher leaves.
We shall see later in what a different way Charles Darwin explained this case of the giraffe. Lamarck's idea is at once clear; it is true that exercising an organ strengthens it, that disuse makes it weaker. Through much gymnastic exercise the muscles of the arm become thicker and more capable, and memory too may be improved, that is to say, even a definite part of the brain may be considerably strengthened by use. Indeed, we may now go so far as to admit that every organ is strengthened by use and weakened by disuse, and so far the foundations of Lamarck's interpretations are sound. But he presupposes something that cannot be admitted so readily, namely, that such 'functional' improvement or diminution in the strength ofan organ can be transmitted by inheritance to the succeeding generation. We shall have to discuss this question in detail at a later stage, and I shall only say now that opinions as to whether this is possible or not are very much divided. I myself doubt this possibility, and therefore cannot admit the validity of the Lamarckian evolutionary principle in so far as it implies the directly transforming effect of the functioning of an organ. But even if we recognize the Lamarckian factor as avera causa, it is easy to show that there are a great many characters which it is not in a position to interpret. Many insects which live upon green leaves are green, and not a few of them possess exactly the shade of green which marks the plant on which they feed; they are thus protected in a certain measure from injuries. But how could this green colour of the skin have been brought about by the activity of the skin, since the colour of the surroundings does not usually stimulate the skin to activity at all? Or how should a grasshopper, which is in the habit of sitting on dry branches of herbs, have thereby been incited to an activity which imparts to it the colour and shape of a dry twig? Just as little, or perhaps still less, can the protective green colour of a bird's or insect's eggs be explained through the direct influence of their usually green surroundings, even if we disregard the fact that the eggs are green when they are laid—that is, before the environment can have had any influence on them.
The Lamarckian principle of modification through use does not, in any case, nearly suffice as an interpretation of the transformations of the organic world. It must be allowed that Lamarck's theory of transformation was well founded at the time when it was advanced; it not only attacked the doctrine of the immutability of species, but sought for the first time to indicate the forces and influences which must be operative in the transformations of species; it was therefore well worth careful testing. Nevertheless it did not divert science from its chosen path; very little notice was taken of it, and in the great Cuvier's chronicle of scientific publications for 1809, not a syllable is devoted to Lamarck's book, so strong was the power of prejudice.
But, although the new doctrine was thus ignored, it did not altogether fall to the ground; it glimmered for a while in Germany, where it found its champions in the 'Naturphilosophie' of the time, and especially in Lorenz Oken, a peasant's son, born at Ortenau, near Offenburg, in 1783.
Oken professed views similar to those of Erasmus Darwin, Treviranus, and Lamarck, though they were not clothed in suchpurely scientific garb, being, in fact, bound up with the general philosophical speculations which came increasingly into favour at that time, chiefly through the writings of Schelling. In the same year, 1809, in which Lamarck published hisPhilosophie zoologique, Oken'sLehrbuch der Naturphilosophieappeared.
This book is by no means simply a theory of descent; its scope is much wider, including the phenomena of the whole cosmos; on the other hand, it goes too little into details and is too indefinite to deserve its title. Its way of playing with ideas, its conjectures and inferences from a fanciful basis, make it difficult for us now to think ourselves into its mode of speculation, but I should like to give some indication of it, for it was just these speculative encroachments of the 'categories' of the so-called 'Naturphilosophie' which played a fatal part in causing the temporary disappearance of the Evolution-theory from science, so that, later on, it had to be established anew.
Oken defines natural science as 'the science of the everlasting transmutations of God (the Spirit) in the world': Every thing, considered in the light of the genetic process of the whole, includes, besides the idea of being, that of not-being, in that it is involved in a higher form. 'In these antitheses the category of polarity is included. The simpler elementary bodies unite into higher forms, which are thus merely repetitions at a potential higher than that of their causes. Thus the different genera of bodies form parallel and corresponding series, the reasonable arrangement of which results as an intrinsic necessity from their genetic connexion. In individuals these lowlier series make their appearance again during development. The contrasts in the solar system between planets and sun are repeated in plants and animals, and, as light is the principle of movement, animals have the power of independent movement in advance of the plants which belong to the earth.'
Obviously enough, this is no longer the study of nature; it is nature-construction from a basis of guesses and analogies rather than of knowledge and facts. Light is the principle of motion, and as animals move, they correspond to the sun, and plants to the planets! Here there is not even a hint of a deepening of knowledge, and all these deductions now seem to us quite worthless.
On the other hand, it must be allowed that good ideas are by no means absent from this 'philosophy,' nor can we deny to this restlessly industrious man a great mind always bent on discovering what was general and essential. Much of what we nowknowhe even then guessed at and taught, as, for instance, that the basis of all forms of life in this infinitely diverse world of organisms was one and thesame substance—'primitive slime,' 'Urschleim' as he called it, or, as we should now say, 'protoplasm.' We can therefore,mutatis mutandis, agree with Oken when he says,'Everything organic has come from slime, and is nothing but diversely organized slime.' Many naturalists of the present day would go further, and agree with Oken when he suggests that 'this primitive slime has arisen in the sea, in the course of the planet's (the earth's) evolution out of inorganic material.'
Thus Oken postulated, as the specific vehicle of life, a primitive substance, in essence at least homogeneous. But he went further, and maintained that his 'Urschleim' assumedthe form of vesicles, of which the various organisms were composed. 'The organic world has as its basis an infinitude of such vesicles.' Who is not at once reminded of the now dominantCell-theory? And, in fact, thirty years later, when the cell was discovered, Oken did claim priority for himself. In so doing, he obviously confused the formulating of a problem with the solving of it; he had, quite rightly, divined that organisms must be built up of very minute concentrations of the primitive substance, but he had never seen a cell, or proved the necessity for its existence, or even attempted to prove it. His vesicle-theory was a pure divination, a prevision of genius, but one which could not directly deepen knowledge; it did not prompt, or even hasten, the discovery of the cell. Here, as throughout in his natural philosophy, Oken built, not from beneath upwards, by first establishing facts and then drawing conclusions from them, but, inversely, he invented ideas and principles, and out of them reconstructed the world. In this he differs essentially from his predecessors Erasmus Darwin, Treviranus, and Lamarck, who all reasoned inductively, that is, from observed data.
Thus the whole evolutionary movement was lost in indefiniteness; because men wanted to find a reason for everything, they missed even what might then have been explained. Moreover, the theory of evolution still lacked a sufficiently broad basis of facts; the 'Naturphilosophie,' by its want of moderation, robbed it of all credit; and it is not to be wondered at that men soon ceased to occupy themselves with the problem of the evolution of the living world. A few indeed held fast to the doctrine of evolution during the first third of the century, but then it disappeared completely from the realm of science.
Its last flicker of life was seen in France, in 1830, at the time of the July revolution, when the legitimate sovereignty of Charles X was overthrown. It is interesting to note the lively interest thatGoethe, the first forerunner of the theory, and then aged eighty-one, had in the intellectual combat that took place in the French Academy between Cuvier and Isidore Geoffroy St.-Hilaire. A friend of Goethe's, Soret, relates that on August 2, 1830, he went into the poet's room, and was greeted with the words: 'Well, what do you think of this great event? The volcano is in eruption, and all is in flames. There can no longer be discussion with closed doors.' Soret replied: 'It is a terrible business! But what else was to be expected with things as they are, and with such a ministry, than that it should end in the expulsion of the reigning family?' To which Goethe answered: 'We don't seem to understand each other, my dear friend. I am not talking of these people at all; I am thinking of quite different affairs. I refer to the open rupture in the Academy between Cuvier and Geoffroy St.-Hilaire; it is of the utmost importance to science.'
In this conflict of opinions, Cuvier opposed Geoffroy's conception of the unity of the plan of structure in all animals, confronting him with the four Cuvierian types, in each of which the plan of structure was altogether different, and strongly insisting on the doctrine of the fixity of species, which he maintained to be the necessary postulate of a scientific natural history.
The victory fell to Cuvier, and it cannot be denied that there was much justification for his opinions at the time, for the knowledge of facts at that stage was not nearly comprehensive enough to give security to the Evolution theory, and moreover the quiet progress of science might have been hindered rather than furthered by premature generalization and theorizing. It had now been seen how far the interpretation of general biological problems could be carried with the available material; the 'Naturphilosophie' had not merely exploited it as far as possible, but had burdened it much beyond its carrying power, and the world was weary of insecure speculations. The 'Naturphilosophie' was for the time quite worked out, and a long period set in, during which all energies were devoted to detailed research.